The Biology of Panchlora nivea, with Observations on the Eggs of

Transcription

The Biology of Panchlora nivea, with Observations on the Eggs of Other Blattaria
Author(s): Louis M. Roth and Edwin R. Willis
Source: Transactions of the American Entomological Society (1890-), Vol. 83, No. 4 (Dec., 1957),
pp. 195-207
Published by: American Entomological Society
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THE BIOLOGY OF PANCHLORA NIVEA, WITH
OBSERVATIONS ON THE EGGS OF
OTHER BLATTARIA
BY LOUIS
Pioneering
M.
ROTH
AND
U.
Research
Division,
and Engineering
Center,
Biology
EDWIN
S. Army
Natick,
of Panchlora
R. WILLIS
Quartermaster
Massachusetts
Research
nivea
of the biology of the pale-green Cuban cockroach,
on
observations
Panchlora
nivea
Various
(Linnaeus).
biological
to
P.
Panchlora
refer
nivea.
different species of
may
purportedly
Rehn and Hebard
(1927) listed seven names that have been used
These workers
stated:
for West
Indian species of Panchlora.
"
cases they relate to
There
is a strong possibility
that in most
Little
is known
the dominant West
Indian species, but their arbi
cubensis,
be warranted."
reference
to that species would
hardly
is a synonym
(1955) has shown that P. cubensis Saussure
Gurney
P.
trary
of P.
Hebard
nivea
(1917) stated that Shelford,
(Linnaeus).
in synonymy under
after examining the types, placed P. virescens
P. nivea.
that
without more ex
Rehn and Hebard
stated
(1927)
on
the
P.
information
viridis
(Fabricius) must be con
plicit
type,
sidered to be an unidentifiable
of
the genus.
However,
species
(=P. nivea) many of the
(1917) identified as P. cubensis
adventives
that had been reported by others as P. viridis. Where
the synonymy
the species as P.
is evident, we have discussed
name
we
it
where
have
retained
is
the
nivea;
not,
assigned by
the original observer.
Hebard
Panchlora
nivea
species of the genus in the
in Mexico,
is
distributed
Central
it
Antilles,
widely
one
it
most
and
northern
South
is
the
fre
America,
America;
eastern
encountered
adventive
in
the
United
States, be
quently
in
to
this
bananas
(Rehn and
ing regularly transported
country
to
P.
is found
Hebard
nivea
1927).
(1917)
Hebard,
According
established
in the United
of Browns
States only in the vicinity
an out-of-doors
it is essentially
form that
ville, Texas;
tropical
Greater
is the dominant
and
can not adapt
itself to artificial
and which,
he
surroundings,
north of the tropical areas
stated, can never become established
of this country.
(195)
TRANS. AMER. ENT.
SOC, LXXX1II.
196
BIOLOGY
OF
PANCHLORA
NIVEA
In Puerto Rico, P. nivea is found in the rotting trunks of palm
or coconut trees where
it feeds on the rotting brown fibers in
are
The nymphs
it tunnels
which
1950).
(Sein, 1923; Wolcott,
and resemble those of Pycnoscelus;
they require 100 days
30 nymphs
the female produces
to mature;
per o?theca
(Sein,
and
P.
is viviparous
nivea
that
Scudder
(1890) reported
1923).
a
that
was
confirmed by Riley
this
(1890, 1891). Riley reported
dissected
He
female of this species gave birth to about 30 young.
o?theca contain
another female and found a semicircular-shaped
ing about 44 eggs. The o?theca consisted of a thin membranous
be
sheath which covered the basal half of the egg mass.
Riley
brown
the
in the genital pouch within
occurred
lieved that hatching
abdomen of the female and that this pouch was so large that the
"
".
development
young could move about and undergo
postnatal
of a female P.
found a museum
Hansen
(1894)
specimen
"
"
from the end of
which had an egg capsule protruding
viridis
to
have
claimed
observed P.
her abdomen.
(1934)
Bayford
"
"
30 young,
to
birth
about
P.
exoleta
viridis
Burm.)
(or
giving
and stated that no o?theca was extruded, nor could he find one
Davis
the female.
after dissecting
(1930) stated that a female
to about 20 young without
birth
P. nivea kept in captivity
gave
an
He
two
males to maturity
reared
; one
egg capsule.
producing
to mature,
the other nearly 8 months.
took a little over 6 months
Shelford
(1906) observed a live specimen of P. nivea which had
an o?theca
the o?theca
from the end of her abdomen;
protruding
partially
"
"
"
"
as
was
compared to the
incomplete
complete
con
in the specimen
observed
Shelford
membrane
by Riley.
the egg mass was so large that it
cluded that in his specimen
the brood sac. Chopard
could not be retained
entirely within
from Riley and Shelford,
(1938), probably taking his information
"
"
and P. nivea
viridis
stated that the o?thecal membrane
of P.
formed and tends to disappear.
is incompletely
The o?thecal membrane
of P. nivea when first laid down covers
It is possible
that Riley's
observation,
nearly all of the eggs.
that the membrane
envelops no more than half of the eggs, was
based on an o?theca
in which the embryos were well developed
;
in this case the increase in the size of the developing
eggs may
have
decreased
the area
covered
by
the o?theca.
Actually,
the
ROTH
AND
197
WILLIS
is so thin and practically
for light
o?theca
colorless
(except
amber colored edges) that it is difficult to see the extent of the
membrane
(Plate XXII,
fig. 9) unless the eggs and o?theca are
(1934)
(Plate XXII,
separated
Bayford
fig. 10). Although
"
"
at parturi
claimed that no o?theca was extruded by P.
viridis
tion, it is possible that he observed the young shortly after they
were born by which
time they may have eaten the o?theca and
membranes,
embryonic
"
ous
cockroaches.
a
common
"
practice
among
ovovivipar
a colony of Panchlora
We have established
nivea from a single
female collected by Edna Roth, May
in a
12, 1956, on bananas
store in Framingham,
After 26 days in the labo
Massachusetts.
ratory, kept at 24? C. and 50 to 60 per cent R. H., this cockroach
Nineteen
gave birth to 60 young.
days after giving birth, the fe
male was observed with an o?theca protruding
beyond her wing
she had completely
retracted the o?theca
tips ;within 55 minutes
into her brood sac. Sixty days after o?theca formation
this fe
male gave birth again.
She died October
12, 1956, after having
lived 153 days in the laboratory ; she fed on banana
skins and
Purina
Dog
o?theca,
tightly
Chow
containing
enveloped
At autopsy we found a J-shaped
53 eggs, in the brood sac. The o?theca was
of the body
by the uterus which filled most
checkers.
cavity.
The
behavior of Panchlora
nivea is similar to that,
ovipositional
in species of Gromphadorhina,
Pycnoscelus,
Nauphoeta,,
and Blaberus
(Roth and Willis,
Leucophaea,
1954). The female
Panchlora
that Shelf ord (1906) reported as carrying an exposed
o?theca was either in the act of ovipositing
or had prematurely
found
the egg case. Premature
extrusion
of the o?theca by
cockroaches
that incubate their eggs in a brood sac may
occur
when they are killed
ex
this
(Roth and Willis,
1954) ;
probably
Hansen's
observation
(1894)
plains
(see above).
to Riley's
Contrary
opinion the nymphs, during normal birth,
do not move about in the genital pouch after the eggs hatch. We
have observed birth once in P. nivea, and the act of parturition
in this species is no different from that in species of Leucophaea,
extruded
Nauphoeta,
re-extruded,
etc.; the o?theca containing
eggs is
fully developed
and once the embryos are clear of the confines of the
TRANS. AMER. ENT.
SOC, LXXXIII.
198
BIOLOGY
OF
PANCHLORA
NIVEA
female's body they free themselves
from their en
completely
and drop from the o?theca.
veloping membranes
Apparently
under abnormal
it is possible
conditions
for an embryo to hatch
one of our females died, she partly
within the brood sac. When
extruded her o?theca.
None of the exposed embryos were able to
free themselves
dissecting
two parts;
this
from the enveloping membranes
and o?theca.
female we found that the o?theca had broken
On
into
the anterior part, containing
ten embryos,
remained
sac.
of
invested
end
the
the
anterior
brood
One em
tightly
by
shed
had
its embryonic
skin almost
bryo
completely;
only its
tarsi were still encased.
This nymph had already become tanned,
and it is conceivable
that at a later stage of development
it might
have shed its skin completely
and moved
about in its cramped
This abortive hatching was possible because the broken
quarters.
left an empty space in the brood sac
o?theca, having separated,
which was prevented
from collapsing
by the eggs that distended
the anterior and posterior
sac contracts to a fraction
An empty brood
parts of the uterus.
of its distended
size.
from the first litter of 60 were reared to
Twenty-three
nymphs
1matured
=
in an average of 144 ? 8.0 x
Males
10)
(N
maturity.
=
The mean number
13) in 181 ? 6.3 days.
days and females (N
of nymphs per litter from 14 births was 46.1 ? 2.4 (range = 28 to
The mean time to the birth of the first litter for seven fe
60).
= 50 to 62
males
isolated after mating was 55.3 ?1.8
days (range
If we allow about a week after mating
to
for oviposition
days).
takes an average of about 48 days at
occur, then embryogenesis
24? C;
it will be recalled, however,
that the first female used to
the P. nivea colony, gave birth 60 days after o?thecal
establish
formation.
Three of the seven females gave birth again 63.3 ?
2.9 days after the first birth.
One female gave birth to a third
litter.
We
have
observed
behavior
the mating
of P. nivea
several
When
confronted with six-day-old
virgin females, males
that are several days old simply run rapidly around behind the
female, back up to her (Plate XXII,
figs. 1, 2), and quickly make
connection.
The final mating
time the
during which
position,
times.
1N
The
= number
at
figures
of
individuals
the right of ?
from which
are standard
mean
errors.
values
were
computed.
ROTH
AND
WILLIS
199
is transferred,
is end to end with the heads of both
spermatophore
directions
individuals
(Plate XXII,
facing in opposite
fig. 3).
=
The average time spent in copulo was 40 ? 2.4 minutes
(range
=
females mated
30 to 46 minutes, N
7). Six-day-old
readily ; the
court
males mated with these without
any apparent preliminary
ter
ship. The female does not feed on a secretion on the male's
gum nor does she assume the superior position prior to copula, as
for several other species
has been reported
(Roth and Willis,
four and five days old did not mate as readily as
1954). Females
those six days old. With
these younger
females, the male be
He moved
the female
around
and
differently.
slowly
near
the
of
her
rocked
end
abdomen
he
from
side
where
stopped
so
to side and at the same time arched his abdomen downwards
that his wings were held slightly away from his body. He then
haved
and
slowly backed up toward the end of the female's abdomen
remained quiescent
in this position
for some time. The younger
female may actively
resist the male's attentions
by kicking with
her hind legs or by running away.
The precopulatory
in Panchlora,
in which
the male
position
to
the female and makes
differs
connection,
simply backs up
from that known
for all other cockroaches
in which mating
has
(1923) stated that the male of
Although Ramme
backed up to the female, this was questioned
Ectobius
by us
Rau
(1957) and found not to be true in Ectobius
pallidus.
on the mating
of Parco
(1922) made a single field observation
blatta pensylvanica
and claimed that the female was
(DeGeer)
been observed.
the aggressor and attempted
to back under the male
(this would
be a male superior pose).
This precopulatory
is highly
position
doubtful.
We have observed
the courtship
of Parcoblatta
vir
like other species of cock
ginica (Brunner) ; this species behaves
roaches in which the males have a tergal gland.
The male of P.
after touching the female with his antennae,
raises his
virginica,
for a
wTings to about a 45? angle ;he may remain in this position
time
if the female
that he is courting does not respond.
If the fe
on
to
feed
his
he
raises
his
begins
tergal secretion,
wings to
and
her
when
reach
his
first
abdominal
90?,
mouthparts
segment,
he pushes backwards
and grasps her genitalia.
The pair then as
sume a position,
of blattids, with their heads facing in
typical
male
TRANS. AMER. ENT.
SOC, LXXXIII.
200
BIOLOGY
PANCHLORA
OF
NIVEA
to court
Males
of P. virginica are attracted
opposite directions.
on
as
and feed
the male
the fe
ing males
just
tergal secretion
this feeding
stimulates
the courting male
to attempt
males;
copulation.
In practically
all species of cockroaches,
the male
just prior to genital connection.
so
far
as we
know,
are
Panchlora
nivea
the
The
and
female
only
Pycnoscelus
is above
exceptions
surina
(Linnaeus) ? In this latter species we did not observe any
he simply climbed onto the back of the
courtship by the male;
female and twisted his abdomen down and under that of the fe
male
from
(Plate XXII,
figs. 4-6) ; connection was readily made
mensis
this male
like the behavior
of grass
superior pose, somewhat
After
connection
has
been
final
the
hoppers.
made,
posi
mating
tion (Plate XXII,
figs. 7, 8) is similar to that of other cockroaches.
was 23 ? 1.2 min
The mean time spent in copulo by Pycnoscelus
=
utes
=15
N
to
26
time the
(range
8), during which
minutes,
was
transferred
to
the
female.
spermatophore
Water
Relations
of
the
Eggs
of
Some
Blattaria
the water uptake by cockroach
(1955) we discussed
Recently
water
and
showed
content
that
the
initial
of eggs and
eggs
a
in
o?thecae of species that incubate their eggs
brood sac is con
lower
water
content
of
than the
the o?thecae and con
siderably
1
of
tained eggs of oviparous
Table
gives the percentage
species.
water not previously
in
and
the newly formed o?thecae
reported
The relationship
of a rela
eggs of seven species of cockroaches.
tively low initial water content in the eggs and o?thecae of false
in comparison
the oviparous
cockroaches
with
ovoviviparous
2
is
forms (except Ectobius
pallidus)
again borne out. Table
shows the changes with age in water and dry matter
content of
nivea and two species of blaberine
cockroaches
eggs of Panchlora
which also incubate their eggs in a brood sac. The water content
of the eggs of these species increased with
increase in dry matter which
corresponding
embryonic
1955a).
development
inDiploptera
age but there was
is so characteristic
punctata
no
of
(Roth and Willis,
2
in the United
and
surinamensis
is parthenogenetic
States
Pycnoscelus
no males
occur
in this country.
in nature
We
obtained
three
apparently
were
in a
reared
which
included
males
from nymphs
that were
accidentally
of Diploptera
from Hawaii.
A bisexual
(Esehseholtz)
shipment
punctata
in our laboratory.
has been
of Pycnoscelus
established
colony
ROTH
WILLIS
201
and
of newly
formed o?thecae
content
from several species of cockroaches
1. Water
Table
AND
enclosed
their
eggs
(%)?S.E.iof
and eggs
o?thecae
and abandon
deposit
Oviparous
species which
after
its formation:
the o?theca
shortly
(Olivier)
(1) Ectobius
pallidus
(2) Neostylopyga
rhombifolia
(2) Byrsotria fumigata (Gu?rin)
nivea
(Linnaeus)
40.0
64.3
56.9
64.5
(Stoll)
(Brunner)
(3) Parcoblatta
virginica
brunnea
Burmeister
(4) Periplaneta
which
extrude
False
species
ovoviviparous
and carry it inter
the o?theca
then retract
development
nally until the eggs complete
Burmeister
(1) Blaberus
craniifer
(3) Panchlora
content
Water
and type of
Species
behavior
oviposition
?
?
?
?
0.61
0.61
0.68
0.43
N:
5
5
5
5
:
1
36.0
34.7 ? 0.56
37.0 1
3
1 S. E. = standard
error
2N = number
of o?thecae
seems to un
in the egg which
the eggs pick up
Sometime
before hatching,
dergo diapause.
water which causes the o?theca to swell (Roth and Willis,
1957).
is known about the rate of uptake of water by the eggs
Nothing
o?thecae
of this species.
On April 16, 1957, some overwintering
and a few nymphs
(older than first instar) of E. pallidus were
Ectobius
pallidus
overwinters
in Plymouth,
These o?thecae had not
Massachusetts.
collected
content of five of these
The water
yet begun to swell visibly.
o?thecae was 41.0 ? 1.1 per cent. We had previously
determined
content of newly formed o?thecae
that the water
of this species
was about 40 per cent (Table 1).
in the laboratory)
(deposited
Thus
water
the
of Ectobius
do not pick up
pallidus
apparently
winter
water
the
content of the
the
months
since
during
eggs collected in the field in April
(and which had been deposited
summer or fall) was essentially
in the previous
the same as that
of newly formed o?thecae.
The o?thecae were divided into two groups ; one group was kept
on filter paper moistened
with distilled water;
the
continuously
eggs
from moist
group was removed
paper after nine days.
of
w^ater
is
shown
in
1A. Both groups of
uptake
text-figure
o?thecae
increased
in
four
for
those eggs
rapidly
weight
days;
moist
solid
then
line)
kept
1A,
continuously
(text-fig.
slowrly in
second
The
creased
in weight
TRANS. AMER. ENT.
from the fourth to the twenty-first
SOC, LXXXIII.
day at which
202
biology
2. Changes
in wet weight, water, and dry matter
cockroaches
of three species of false ovoviviparous
Table
:The
Note
o?thecae,
weight
age.
were
nivea
panchlora
of
actual
were
ages
of
the
or recently
formed
for the newly
except
are arranged
in order of increasing
wet
an increase
in
with
development,
represents
eggs,
The
unknown.
eggs
correlated
which,
being
in the computations.
is included
The weight
of the o?theca
determined
for each wet weight
listed.
from one o?theca
Milligrams
Species
Blaberus
Wet
Dry
weight
13.21
craniifer
19.8
25.4
25.6
30.02
Byrsotria
fumigata
13.11
15.3
8.6
4.5
!
0.41
0.3
Per Egg
Matter
Water
8.4 4.8
7.6 12.2
7.1 18.3
7.4 18.2
8.2 21.8
1.02
8.8
9.8
7.8
9.4
18.6
18.1
20.6
20.0
0.1
0.7
0.8
1
or recently
formed
Newly
2
Well-developed
embryos.
values
10.6 5.4
27.4
27.9
28.4
29.4
nivea
All
9.85.5
16.0 !
Panchlora
of eggs
0.2 0.5
0.2 0.6
0.3 0.7
o?thecae.
time the eggs began to hatch.
It is estimated
from known dry
of o?thecae that at the time of hatching the o?thecae and
weights
The second
eggs of group one contained about 75 per cent water.
increased
group of o?thecae
(text-fig.
1A, broken line) similarly
in weight until the ninth day. After the ninth day the eggs were
placed on dry filter paper and they began to lose water gradually.
in spite of this water
loss the eggs in these o?thecae
However,
hatched on the twenty-second
and twenty-third
It is ap
days.
if sufficient moisture
is absorbed, the eggs may subse
some
lose
when
to dry conditions,
but
water,
subjected
quently
still complete development
and hatch.
parent
that
I
I
I
I
I
I
ROTH
AND WILLIS
I
I
I
I
I
I
fe 60F
?C Ul
O O
203
I
I
I
I
I
I
I
I
^-.^
^_
< 40
?J
or
o
z20h
.H
z
?
I
0
?
i i i i i i
4
0
16
12
8
20
24
32
36
pallidus.
into the
labora
28
DAYS
of
Uptake
overwintered
1.?A.
Text-figure
o?thecae
had
tory on 16 April,
were
the o?thecae
water.
The
= Six
.-.
on
to hatch
.-.
days;
see
The
axis
of abscissas
on filter paper
placed
are mean
values.
points
kept
on moistened
the
represents
moistened
paper
until
with
the
The
days after
distilled
eggs
began
the
day.
twenty-first
on moistened
o?thecae
kept
on dry filter
One
o?theca
kept
of Ectobius
eggs
by
in the field and were
taken
first
plotted
o?thecae
= Five
then
.=
1957.
water
paper
for
filter
paper
nine
kept
on
moistened
text.
TRANS. AMER. ENT.
days
and
paper.
SOC., LXXXIII.
for
fourteen
40
204
B.
of
O?thecae
axis
The
of
biology
is expressed
weight
was
last weighing
(H)
.-.
on
the
= Ten
relative
the
Parcoblatta
abscissas
represents
are mean
values.
= Five
o?thecae
points
.-.
as
made
thirty-ninth
o?thecae
virginica
the age
nivea
in
deposited
of the o?thecae.
on moistened
kept
the per cent
increase
on
the
thirty-sixth
filter
of
day
paper
the
the
; the
initial
and
laboratory.
The
plotted
the
in
change
The
weight.
eggs
hatched
day.
humidity.
The
kept
doss
weight.
None
of
initial
panchlora
of
on
at
paper
dry filter
in weight
is expressed
the eggs hatched.
50 to 60 per
as a percentage
A single o?theca behaved somewhat differently
from the
are
its weight
changes
plotted
1A,
separately
(text-fig.
line). This o?theca showed very little increase in weight
first three days on moist filter paper, but after the third
cent
of
others
;
dotted
for the
day it
more
to
water
at the time
began
pick up
rapidly.
Apparently
this o?theca was collected,
the eggs had not reached the stage in
to absorb water
wThen they begin
the
development
through
o?theca.
By the fourth day water began to be absorbed, and the
curve is similar to the others described previ
subsequent weight
After the twelfth day on moist paper, this o?theca began
ously.
so it wTas dissected on the fourteenth day; many of
to lose weight,
the eggs were well developed
having dark eye spots, but some
had died and therefore
had begun to lose
embryos apparently
weight.
In the field, the eggs of Parcoblatta
are deposited
in
virginica
under logs, bark, etc. In the labora
relatively moist
situations,
tory, the female usually deposits her o?thecae on the moist cotton
stoppers of vials containing
drinking wrater. Those eggs not de
cotton do not develop at room humidities
posited on the moist
and the o?thecae
This is similar to the be
collapse.
frequently
havior of the eggs of Cariblatta
lutea minima Hebard.
The initial water content of o?thecae and eggs of P. virginica
is about 57 per cent (Table 1). After two weeks, when kept on
moist
o?theca
filter
and
paper, the eggs begin to absorb water
through the
continue to do so at a slow uniform rate, as repre
sented
occurs
IB, solid line), until hatching
by weight
gain (text-fig.
on the thirty-ninth
It
is estimated
that at the time
day.
of hatching
the o?thecae
and eggs contained
about 67 per cent
water.
If the eggs are kept on dry filter paper at 50 to 60 per
AND
ROTH
205
WILLIS
cent R. H., water
is lost steadily, as represented
loss
by weight
(text-fig. IB, broken line), and the eggs fail to develop.
of P. virginica
differ from those of some other
The o?thecae
americana)
oviparous
species (e.g., Blatta
orientalis, Periplaneta
re
an impermeable
not possessing
in apparently
coating which
that are exposed to relatively
tards loss of water
from o?thecae
do not undergo
The eggs of P. virginica
dia
dry conditions'.
in this respect they apparently
differ from the eggs of
pause;
Ectobius
pallidus.
Summary
Panchlora
nivea
extrudes
its o?theca
formation
and
during
When
then retracts it into a brood sac where the eggs develop.
the embryos mature,
the nymphs
free
the o?theca is re-extruded;
as the o?theca
is
themselves
from their enveloping membranes,
forced from the brood sac, and drop from the female.
One mat
ing enabled a female to produce three litters. The mean number
of nymphs per litter was 46. Embryogenesis
took about 48 days
at 24? C. Male nymphs matured
in an average of 144 days and
females
in 181 days.
The mating
has been observed
from that which
behavior
of P.
in other
nivea
differs
in that
cockroaches
the male backs up to the female to make genital connection,
and
the female does not feed on any secretion on the male's
tergum.
behavior among the Blattaria
is that of
Another peculiar mating
the male of which makes
connection
Pycnoscelus
surinamensis,
with the female after climbing on her back.
The final mating
and Pycnoscelus
is similar to that of
position of both Panchlora
other cockroaches;
the sexes, with genitalia
joined, face in oppo
site directions.
re
The water
contents
of o?thecae
and eggs, not previously
are given.
The eggs of
ported for several species of cockroaches
Panchlora
nivea,
crease in water
Blaberus
in
craniifer, and Byrsotria
fumigata
but not dry matter,
content,
during embryo
The eggs of Ectobius
overwinter
in the field
genesis.
pallidus
no change in water content.
with essentially
of E. pal
O?thecae
lidus that had overwintered
in nature were brought into the labo
ratory and kept moistened
days and then increased
TRANS. AMER. ENT.
; they absorbed water
in weight
slowly until
rapidly for four
the twenty-first
SOC, LXXXIII.
206
BIOLOGY
OF
PANCHLORA
NIVEA
of Parcoblatta
vir
the eggs hatched.
The o?thecae
day when
are
In
in
the
moist
situations.
eggs
ginica
laboratory,
deposited
of P. virginica
that were kept moist began to absorb water after
on the
two weeks and increased in weight
slowly until hatching
were
P.
When
the
o?thecae
of
virginica
kept at
thirty-ninth
day.
50 to 60 per cent relative
and did not hatch.
lost water
the eggs
humidity,
steadily
Acknowledgments
We
thank Dr.
Barbara
the specimens
and
Stay for preparing
the photographs
for figures 9 and 10, and for collecting,
of Ectobius
together with Roger Swain, the o?thecae
pallidus;
and Dr. Ashley B. Gurney, United
States National
for
Museum,
our specimens of Panchlora
nivea.
identifying
taking
References
E.
in cockroaches.
Ent.
Monthly
Mag.
Viviparity
: 210.
[3rd. Ser., vol. 20(237)]
70(844)
1938.
La biologie
des Orthopt?res.
S?r. A, 20:
ARD, L.
Encycl.
Ent.,
1-541.
Bayford,
Chop
Davis,
T.
W.
1934.
G.
1930.
lora cubensis.
the young
Rearing
Jour.
New
Notes
on
Gurney,
A. B.
Hansen,
Blattidae).
(Orthoptera,
H. J.
1894.
On the
1955.
Walk.
Ent.
Hebard,
M.
Ramme,
boundary.
1923.
W.
1917.
Ectobius
Rau,
P.
1922.
Acad.
Rehn,
York
of
Ent.
the Cuban
Proc.
Ent.
structure
Amer.
Vorarbeiten
Arch.
Steph.
and M.
1.
Hebard.
Blattidae.
habits
nivea
57:
of Hemimerus
of
north
America
Panch
(L.)
285-286.
talpoides
the Mexican
Ent.
Soc, No.
2, 284 pp.
zu einer Monographie
des Blattidengenus
f. Naturges.,
Abt. A, 89: 97-145.
and behavior
Ecological
24: 1-71.
Sei. St. Louis,
J. A. G.,
Number
Panchlora
cockroach,
Soc. Washington,
and
15: 65-93.
Tidsk.,
The Blattidae
of North
Mem.
the viviparous
cockroach
38: 85-88.
Soc,
1927.
Bull.
on Missouri
notes
The
Amer.
insects.
of the West
Orthoptera
Nat.
Mus.
54
Hist.,
Trans.
Indies.
(Art.
1):
1-320.
Riley,
C. V.
2:
-
Roth,
1890.
A
viviparous
cockroach.
Proc.
Ent.
Soc. Washington,
129-130.
1891.
Further
notes
on Panchlora.
Insect
and E. R. Willis.
L. M.,
1954.
The
Smithson.
122 (12) : 1-49.
Misc.
Coll.,
1955. Water
content
of cockroach
in relation
to oviposition
behavior.
Jour.
eggs
Exp.
4:
Life,
of
reproduction
during
Zool.,
119-120.
cockroaches.
embryogenesis
128: 489-509.
ROTH AND WILLIS
-
-
(Serv.)
dytiscoides
in the laboratory
Se?n,
F.,
Shelford,
Jr.
1923.
G.
of
Studies
Trans.
Ent.
1950.
N.
Puerto
Amer.
Rico,
62:
Psyche,
pallidus
Diploptera
its biology
(Oliver)
Stat.
55-68.
(Blattaria,
Circ,
No.
of
the
VI. Viviparity
amongst
: 509-514.
(1906)
Puerto
Rico.
Jour. Agr.
Univ.
of
Figures
London
insects
32:
on
notes
with
the Blattidae.
Soc.
The
1948,
"
beetle-roach
83: 31-37.
Ent.
Soc,
5: 405.
Psyche,
Puerto
Insular Exp.
Rico
title].
[No
"
Diplopteridae).
of Ectobius
Cucarachas.
64, 12 pp.
1906.
R.
Blattidae.
Wolcott,
(Blattaria:
Trans.
1890.
S. H.
the
embryogenesis,
during
1957. The biology
Blattidae).
Scudder,
of
nutrition
Intra-uterine
1955a.
207
1-224.
Explanation
PLATE XXII
Figs.
female.
1-3.?Panchlora
3. Pair
in copulo.
2. Male
nivea.
(blurred)
Vertical
Figs.
4-8.?Pycnoscelus
to make
connection.
sume
around
moving
female
and
surinamensis.
4-6. Male
on
to posterior
end of
connection.
making
female
7. Pair
with male
connected,
twisting
8. Final mating
position.
nivea.
9. O?theca
and eggs removed
an upright
position.
9-10.?Panchlora
Figs.
10. Thin
membrane
o?thecal
bar = 3 mm.
TRANS. AMER. ENT.
1. Male
into
backing
bar = 1 cm.
separated
from
the
enclosed
eggs.
SOC, LXXXIII.
attempting
to as
around
from
female.
Horizontal
Trans.
Amer.
Ent.
Soc,
Vol.
Plate
LXXXIII.
ROTH AND WILLIS:
PANCHLORA
NIVEA
XXII.

The Biology of Panchlora nivea, with Observations on the Eggs of

Transcription

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